Boraginaceae Juss.

Boraginaceae, H. Heine. Flora of West Tropical Africa 2. 1963

Morphology General Habit

Trees, shrubs or herbs, often roughly scabrid or hispid

Morphology Leaves

Leaves alternate or rarely opposite, simple; stipules absent

Morphology Reproductive morphology Flowers

Flowers often in scorpoid cymes, actinomorphic or rarely oblique, mostly hermaphrodite

Morphology Reproductive morphology Flowers Calyx

Calyx-lobes imbricate or rarely valvate

Morphology Reproductive morphology Flowers Corolla

Corolla with contorted or imbricate lobes

Morphology Reproductive morphology Flowers Androecium

Anthers 2-celled, opening lengthwise Stamens the same number as the corolla-lobes and alternate with them, inserted on the corolla

Morphology Reproductive morphology Flowers Nectaries

Disk present or absent

Morphology Reproductive morphology Flowers Gynoecium

Ovules paired, erect or spreading from the central axis Ovary superior, 2-celled or 4-celled by spurious septa, entire or deeply 4-lobed; style terminal or gynobasic

Morphology Reproductive morphology Fruits

Fruit a drupe or of 4: nutlets

Morphology Reproductive morphology Seeds

Seeds with or without endosperm

Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://creativecommons.org/licenses/by/3.0

Morphology General Habit

Herbs, subshrubs, shrubs, trees, rarely lianas; rarely obligate root parasites with chlorophyll absent in Lennoa Lex

Morphology General Hair

Hairs usually present, occasionally containing cystoliths, simple or glandular

Morphology Leaves

Leaves usually alternate, rarely subopposite (Tournefortia L.), sometimes basal (Plagiobothrys Fisch. & C.A.Mey.), simple, rarely compound (Phacelia Juss.), reduced to scales in Lennoa, exstipulate; lamina entire, usually rough to hispid, margins mostly entire, or crenate, or serrate, or dentate with venation pinnate ;  domatia occurring in Cordia L. and Ehretia P.Browne.

Morphology Reproductive morphology Inflorescences

Inflorescence terminal, axillary, or intermodal, cymose, the branches often scorpioid, helicoid, or reduced and capitate to glomerate (as Varronia P.Browne), rarely reduced to one solitary flower (some Heliotropium L.)

Morphology Reproductive morphology Flowers

Flowers actinomorphic, sometimes distylous (Varronia and Cordia), usually 5- merous, hermaphrodite or unisexual; calyx usually 5- lobed, distinct or connate

Morphology Reproductive morphology Flowers Corolla

Corolla sympetalous, usually 5- lobed; stamens isomerous, alternate with corolla lobes; filaments adnate to  corolla tube; anthers 2-locular, dehiscing longitudinally;  ovary superior, 2-carpellate, but usually becoming falsely 4-locular, with axile placentation or often parietal with 2 placentae in Phacelia; ovules usually 4, anatropous to hemitropous; the style terminal or gynobasic, simple, or bifid (Ehretioideae, Phacelia), or twice 2- cleft (Cordia and Varronia); stigmas 1-4;  nectar disk usually present around base of ovary

Morphology Reproductive morphology Fruits

Fruits fleshy or dry drupaceous, or mericarps 2 (2-seeded) or 4 (4-seeded), or a capsule ; seeds 4 per fruit, often less by abortion; endosperm present or absent; cotyledons entire of flat, or rarely plicate (Cordia and Varronia).

Diagnostic

Cystoliths commonly present. Distinguishing characters (always present): Leaves usually alternate, exstipulate. Inflorescence cymose, the branches often scorpioid, helicoid, or reduced and capitate to glomerate. Corolla gamopetalous. Stamens isomerous inserted in the corolla tube. Ovarysuperior, 2-carpellate, but usually becoming falsely 4-locular. Habit Inflorescence Style Fruit Notable genera and distinguishing features: Cordia is characterized by the habit (trees or shrubs); inflorescence cymose, the branches often scorpioid, helicoid; terminalstyle twice bifid forming four stigmatic lobes; fruitdrupaceous.  Varronia is characterized by the habit (multi-stemmed shrubs or rarely several-stemmed treelets); inflorescence reduced, capitate, glomerate, spicate, or if cymose, these less than 4 cm broad; terminalstyle twice bifid forming four stigmatic lobes; fruitdrupaceous.  Heliotropium is characterized by a terminalstyle and dry fruits, which divide into four or two mericarps. Tournefortia is characterized by a terminalstyle and drupaceous fruits, which never divide into mericarps. Ehretia is characterized by the habit (trees or shrubs), and bifid terminalstyle.

Note

Number of genera: The Boraginaceae comprise approximately 205 genera and 2500 species worldwide, of which 23 genera and approximately 500 species are neotropical.   Neotropical genera: Amsinckia Lehm. Antiphytum DC. ex Meisn. Bourreria P.Browne Cordia (including Auxemma Miers, Patagonula L.and Saccellium Humb. & Bonpl.) Cryptantha Lehm. ex Fisch. & C.A.Mey. Ehretia Hackelia Opiz Heliotropium Lasiarrhenum I.M. Johnst. Lennoa Lepidocordia Ducke Lithospermum L. Macromeria D.Don Moritzia DC. ex Meisn. Nama L. Pectocarya DC. ex Meisn. Phacelia Plagiobothrys Rochefortia Sw. Tiquilia Pers. Tournefortia Wigandia Kunth Varronia P.Browne Notes on delimitation: The family is circumscribed broadly, including Hydrophyllaceae (except Hydrolea L.) and Lennoaceae. It is considered monophyletic on the basis of inflorescence type and DNA sequences data (Ferguson 1999; Gottschling et al. 2001).

Distribution

Many species have been introduced as ornamental plants or weeds, e.g. the genera Anchusa L., Borago L., Cynoglossum L., Echium L., Myosotis L. and Symphytum L.. The genus Varronia is endemic.

Boraginaceae, B. Verdcourt. Flora of Tropical East Africa. 1991

Morphology General Habit

Annual or perennial herbs, often woody at the base, subshrubs, trees or woody climbers, usually characteristically scabrid or hispid

Morphology Leaves

Leaves simple, alternate or less often opposite, petiolate or sessile, entire or variously toothed or crenate; stipules absent

Morphology Reproductive morphology Inflorescences

Inflorescences terminal or axillary, bifurcate or in raceme-like or panicled groups, basically dichotomous cymes, the branches frequently scorpioid and spirally inrolled when young; bracts present or absent

Morphology Reproductive morphology Flowers

Flowers hermaphrodite or unisexual (Cordia), sessile or pedicellate, mostly 5-merous, regular or somewhat irregular, occasionally heterostylous

Morphology Reproductive morphology Flowers Calyx

Calyx tubular or campanulate, persistent and sometimes accrescent in fruit, with (2–)4–7 (or more) imbricate or rarely valvate or open lobes

Morphology Reproductive morphology Flowers Corolla

Corolla tubular, campanulate, funnel-shaped or rotate with tube very reduced; lobes 3–16, imbricate or contorted (>i>Myosotis); throat often with scales, thickenings, folds or crests

Morphology Reproductive morphology Flowers Androecium

Stamens the same number and alternating with the corolla-lobes, exserted or included, epipetalous; anthers 2- thecous

Morphology Reproductive morphology Flowers Nectaries

Nectariferous disc usually present

Morphology Reproductive morphology Flowers Gynoecium

Style usually 1, 2 in one genus or sometimes 4 in one species of >i>Cordia, terminal or gynobasic, entire or 2–4-fid or twice 2-fid; stigmas clavate, capitate or peltate or linear to narrowly foliaceous Ovules axile, anatropous, erect, horizontal, basal or rarely pendulous Ovary superior, entire or deeply 4-lobed, 2-locular with 2 ovules in each or with 4, 1-ovuled locules due to the development of false partitions

Morphology Reproductive morphology Fruits

Fruit drupaceous with 1, 4-celled endocarp or separating in 2 bilocular or 4 unilocular 1-seeded pyrenes or with 4 quite distinct nutlets

Morphology Reproductive morphology Seeds

Seed with or without endosperm, erect, oblique or horizontal; testa membranous; embryo straight or curved; cotyledons flat or plicate

Boraginaceae, E. S. Martins (Trichodesma by R. K. Brummit). Flora Zambesiaca 7:4. 1990

Morphology General Habit

Herbs, shrubs or rarely trees

Morphology Leaves

Leaves alternate, rarely subopposite or opposite, simple exstipulate

Morphology Reproductive morphology Inflorescences

Inflorescence usually cymose, composed of one or more helicoid or scorpioid cymes, sometimes paniculate or racemose or sometimes flowers solitary, terminal or axillary

Morphology Reproductive morphology Flowers

Flowers hermaphrodite or sometimes unisexual, actinomorphic or sometimes zygomorphic

Morphology Reproductive morphology Flowers Calyx

Calyx (4)5(6)-lobed or 3–5-toothed, imbricate or valvate

Morphology Reproductive morphology Flowers Corolla

Corolla (4)5(6)-lobed, imbricate or contorted in bud, tubular, funnel-shaped, campanulate, salver-shaped or rotate; tube sometimes with folds, scales or hairs in the throat

Morphology Reproductive morphology Flowers Androecium

Stamens (or staminodes) as many as corolla lobes and alternate with them, inserted on the corolla tube; anthers 2-thecous, dehiscing longitudinally, introrse, dorsifixed

Morphology Reproductive morphology Flowers Nectaries

Disk present or absent

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, entire or deeply 4-lobed, bi-locular or spuriously 4-locular, placentation axial, ovules 4, erect or spreading; style 1, terminal or gynobasic, entire or cleft or twice cleft, or styles 2, terminal (Coldenia)

Morphology Reproductive morphology Fruits

Fruit drupaceous with a 1–4-seeded stone or splitting into 4 pyrenes, or of one to four-seeded nutlets; seeds generally without endosperm

Gemma Bramley, Anna Trias-Blasi & Richard Wilford (2023). The Kew Temperate Plant Families Identification Handbook. Kew Publishing Royal Botanic Gardens, Kew

Recognition

Characters of similar families: Heliotropiaceae: style terminal, fruit nutlets or drupe. Hydrophyllaceae: style terminal, fruit a capsule. Lamiaceae: leaves opposite, usually zygomorphic corolla. Solanaceae: fruit many-seeded.

Morphology General Habit

Herbs rarely shrubs; stipules absent

Morphology Leaves

Leaves alternate, simple; lamina often rough to hispid, hairs often with swollen bases, margins entire

Morphology Reproductive morphology Inflorescences

Inflorescence terminal or axillary, usually scorpioid, monochasial or dichasial, occasionally solitary, more often in lax to condensed cymes

Morphology Reproductive morphology Flowers

Flowers blue, white, yellow or pink, often changing colour as they mature, usually actinomorphic, bisexual, usually 5-merous; calyx lobes almost entirely fused to united at base only; corolla sympetalous, 5-merous, tubular, campanulate, salver-shaped, funnel-form or rotate; stamens typically 5, attached to the corolla tube; anthers exserted or not; ovary superior with 2 carpels, 4-locular; style gynobasic, stigma bilobed to capitate

Morphology Reproductive morphology Fruits

Fruit (1–2)4 nutlets, smooth, winged or often ornamented with glochidiate spines

Morphology Reproductive morphology Seeds

Seed with or without endosperm.

Distribution

A family of 94 genera and ca.1,800 species; Boraginaceae is presented here sensu stricto (several large genera are now excluded — Heliotropium (Heliotropiaceae), Phacelia (Hydrophyllaceae)). The family is predominantly northern temperate but extends to mountainous areas of the tropics. Major genera are Cynoglossum, Cryptantha and Myosotis.

Note

Leaves simple, alternate, often rough hairy. Inflorescences cymose, often scorpioid. Flowers actinomorphic, 5-merous, sympetalous; style gynobasic; ovary superior. Fruit a schizocarp, (1–2)4 nutlets.

Description Author

Gemma Bramley

George R. Proctor (2012). Flora of the Cayman Isands (Second Edition). Royal Botanic Gardens, Kew

Morphology General Habit

Annual or perennial herbs, or else shrubs, trees or woody vines, usually with scabrid, setose, or hispid indument, rarely tomentose or glabrous; leaves alternate (rarely opposite or whorled), simple, entire or toothed; stipules none

Morphology Reproductive morphology Flowers

Flowers usually regular, perfect or rarely polygamous by abortion, borne in usually dichotomous cymose spikes, racemes, heads or panicles

Morphology Reproductive morphology Flowers Calyx

Calyx usually 5-toothed or -lobed, more or less bell-shaped, imbricate or often open in bud, usually persistent and often enlarging in fruit

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, usually 5-lobed (rarely 6–8-lobed), tubular, funnel-, bell- or salver-shaped or subrotate, the lobes imbricate or twisted in bud, sometimes with pleats or appendages in the tube or partly closing the throat

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens as many as the corolla-lobes and alternating with them, inserted in the corolla-throat or -tube; filaments free; anthers 2- locular, opening lengthwise, attached above the base and more or less 2-lobed below the point of attachment

Disc

Disc ring-like, entire or 5-lobed, or apparently absent. Ovary superior, 2-locular and 2-carpellate, but often becoming falsely 4-locular at maturity, simple or deeply 4-lobed; ovules usually 4, 2 in each carpel; styles 1 or 2, terminal on simple ovaries, or sometimes arising from between the lobes on lobate ones, simple or 1–2-forked at the apex

Morphology Reproductive morphology Fruits

Fruit of 4 1-seeded nutlets or a 1–4-seeded nut or drupe; seeds with or without endosperm.

Distribution

A cosmopolitan family of about 100 genera and 2,000 species.

Hoplestigmataceae, F.N. Hepper. Flora of West Tropical Africa 2. 1963

Morphology General Habit

Trees

Morphology Leaves

Leaves alternate, entire; stipules absent

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, in terminal cymes

Morphology Reproductive morphology Flowers Calyx

Calyx closed in bud, bursting irregularly

Morphology Reproductive morphology Flowers Corolla

Corolla shortly tubular, with about 3 series of overlapping corolla lobes

Morphology Reproductive morphology Flowers Androecium

Stamens numerous (about 21), in several rows inserted at the base of the corolla; anthers 4-celled, opening by longitudinal slits

Morphology Reproductive morphology Flowers Gynoecium

Ovules pendulous, one on each side of the bilobed placentas Ovary superior, 1-celled with 2 intrusive parietal placentas; style bilobed nearly to the base, branches sharply bent; stigmas capitate

Morphology Reproductive morphology Fruits

Fruit with a deep groove up each side, the calyx forming a plate at the base, endocarp hard

Morphology Reproductive morphology Seeds

Seeds with scanty endosperm and long thick straight embryo

Hydrophyllaceae, F.N. Hepper. Flora of West Tropical Africa 2. 1963

Morphology General Habit

Annual or perennial herbs

Morphology Leaves

Leaves radical or alternate, rarely opposite, entire to pinnately or palmately lobed

Morphology Reproductive morphology Flowers

Flowers usually cymose, hermaphrodite, actinomorphic

Morphology Reproductive morphology Flowers Calyx

Calyx-segments 5, imbricate

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, lobes imbricate or contorted

Morphology Reproductive morphology Flowers Androecium

Stamens the same number as the corolla-lobes and alternate with them, mostly inserted towards the base of the tube; anthers 2-celled

Morphology Reproductive morphology Flowers Gynoecium

Ovules numerous Ovary superior, 1-celled with 2 parietal placentas, or 2-3-celled; styles 1 or 2

Morphology Reproductive morphology Fruits

Fruit a loculicidal or rarely septicidal capsule

Morphology Reproductive morphology Seeds

Seeds with fleshy endosperm and small straight embryo

Hydrophyllaceae, B. Verdcourt. Flora of Tropical East Africa. 1989

Morphology General Habit

Annual or perennial herbs, rarely subshrubs, often scabrid

Morphology Leaves

Leaves usually alternate or radical, exstipulate, entire to deeply pinnately or palmately lobed, the margins entire, serrate or lobulate

Morphology Reproductive morphology Flowers

Flowers regular, hermaphrodite, in scorpioid or trichotomous cymes, false racemes or clustered, less often solitary

Morphology Reproductive morphology Flowers Calyx

Calyx deeply divided into 5 (rarely 8–12) imbricate or open, equal or unequal lobes or 5 with appendages between

Morphology Reproductive morphology Flowers Corolla

Corolla often blue, variously shaped, the lobes usually 5 (rarely 8–12), imbricate or rarely contorted

Morphology Reproductive morphology Flowers Androecium

Stamens as many as the corolla-lobes, inserted at the sinuses or low in the tube, filaments filiform or dilated at the base; anthers 2-thecous, versatile, opening by longitudinal slits, included or exserted

Morphology Reproductive morphology Flowers Nectaries

Disk inconspicuous or absent

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior (very unusually inferior), 1-locular with 2 parietal fleshy placentas or 2(–3)-locular; ovules 2 or more on each placenta; style filiform, usually bifid or styles 2; stigmas simple or capitate

Morphology Reproductive morphology Fruits

Fruit usually a loculicidal or less often septicidal 2-valved or irregularly splitting capsule

Morphology Reproductive morphology Seeds

Seeds oblong, globose or angled, with tuberculate, reticulate or rugose testa; albumen copious and fleshy; embryo small; cotyledons entire

George R. Proctor (2012). Flora of the Cayman Isands (Second Edition). Royal Botanic Gardens, Kew

Morphology General Habit

Annual or perennial herbs, rarely shrubs, often hairy or scabrid, sometimes armed with spines; leaves alternate or rarely opposite, sometimes in basal rosettes, entire to pinnately or palmately divided; stipules absent

Morphology Reproductive morphology Flowers

Flowers perfect and regular, borne in dichasial, umbel-like, or helicoid cymes, or sometimes solitary in the leaf-axils

Morphology Reproductive morphology Flowers Calyx

Calyx 5-lobed, imbricate, the lobes often with appendages between; corolla gamopetalous, 5-lobed, the lobes imbricate or contorted

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens usually 5, inserted on the corolla-tube near its base and alternate with the lobes; filaments often dilated at the base, or subtended by appendages; anthers 2-celled, opening lengthwise

Morphology Reproductive morphology Flowers Gynoecium Ovary

Ovary superior or partly inferior, 2- carpellate but usually 1-celled with 2 parietal placentas meeting in the centre, the ovules 4–numerous; style 1 or 2 (rarely more)

Morphology Reproductive morphology Fruits

Fruit a capsule, dehiscent or rarely indehiscent; seeds variously sculptured; endosperm copious or thin.

Distribution

A widespread family of 20 genera and about 270 species, best represented in N. America, absent from Australia.

Hydrophyllaceae, B. Verdcourt. Flora Zambesiaca 7:4. 1990

Morphology General Habit

Annual or perennial herbs, rarely subshrubs, often scabrid

Morphology Leaves

Leaves usually alternate or radical, exstipulate, entire to deeply pinnately or palmately lobed, with margins entire, serrate or lobulate

Morphology Reproductive morphology Flowers

Flowers regular, hermaphrodite, in scorpioid or trichotomous cymes, false racemes or clustered, less often solitary

Morphology Reproductive morphology Flowers Calyx

Calyx deeply divided into 5 rarely 8–12 imbricate or open equal or unequal lobes or 5 with appendages between

Morphology Reproductive morphology Flowers Corolla

Corolla often blue, variously shaped, the lobes usually 5, rarely 8–12, imbricate or rarely contorted

Morphology Reproductive morphology Flowers Androecium

Stamens as many as the corolla lobes, inserted at the sinuses or low in the tube; filaments filiform or dilated at the base; anthers bilocular, versatile, opening by longitudinal slits, included or exserted

Morphology Reproductive morphology Flowers Nectaries

Disk inconspicuous or absent

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior (very unusually inferior) unilocular with 2 parietal fleshy placentas or 2(3)-locular Ovules 2 or more on each placenta; style filiform, usually bifid or styles 2; stigmas simple or capitate

Morphology Reproductive morphology Fruits

Fruit usually a loculicidal or less often septicidal bivalved or irregularly splitting capsule

Morphology Reproductive morphology Seeds

Seeds oblong, globose or angled with tuberculate reticulate or rugose testa; albumen copious and fleshy; embryo small; cotyledons entire